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Cerithioidea: Thiaridae and Pachychilidae) Systematics, phylogeny and biogeography of Mesoamerican and Caribbean freshwater gastropods (Cerithioidea: Thiaridae and Pachychilidae) D i s s e r t a t i o n zur Erlangung des akademischen Grades d o c t o r r e r u m n a t u r a l i u m ( Dr. rer. nat.) im Fach Biologie eingereicht an der Mathematisch-Naturwissenschaftlichen Fakultät I der Humboldt-Universität zu Berlin von M.Sc. Biologin Maria I. Gómez geb. 09.12.1973 in Pereira (Kolumbien) Präsident/Präsidentin der Humboldt-Universität zu Berlin Prof. Dr. Dr. h.c. Christoph Markschies Dekan der Mathematisch-Naturwissenschaftlichen Fakultät I Prof. Dr. Lutz-Helmut Schön Gutachter/innen: 1. Prof. Dr. Hannelore Hoch 2. PD Dr. Mark-Oliver Rödel 3. Prof. Dr. Thomas Wilke Tag der mündlichen Prüfung: 16.11.2009 “It is perhaps a more fortunate destiny to have a taste for collecting shells than to be born a millionaire. Although neither is to be despised, it is always better policy to learn an interest than to make a thousand pounds; for the money will soon be spent, or perhaps you may feel no joy in spending it; but the interest remains imperishable and ever new” Robert Louis Stevenson Gedruckt mit Unterstützung des Deutschen Akademischen Austauschdienstes. Zusammenfassung In dieser Arbeit habe ich die Systematik zweier distinkter Taxa von Süßwasserschnecken innerhalb der Superfamilie Cerithioidea untersucht: der Familie Pachychilidae Troschel, 1858 und der Gattung Hemisinus Swainson, 1840 aus der Familie Thiaridae Gill, 1871. Durch vergleichende Analysen der morphologischen, anatomischen und molekulargenetischen Daten habe ich den Status der beschriebenen Arten für die gesamten Pachychilidae aus der Karibik und Mittelamerika, sowie für Hemisinus aus dem karibischen Raum überprüft. Des weiteren habe ich die Verwandtschaftsbeziehungen dieser Familien zueinander und zu ihren Verwandten aus anderen tropischen Ländern bestimmt. Anhand von Informationen und Material aus Museumssammlungen und von Geländeuntersuchungen habe ich außerdem die Verbreitung der Vertreter beider Familien rekonstruiert. Innerhalb der neotropischen Thiaridae habe ich alle Hemisinus-Arten erneut und erweitert beschrieben. Die Gattung kommt in der Karibik lediglich auf den Großen Antillen vor und wird durch eine endemische Art auf Jamaika (Hemisinus lineolatus), welche gleichzeitig der Typus ist, sowie vermutlich zwei endemische Arten auf Kuba (H. cubanianus und Cubaedomus brevis) vertreten. Morphologisch unterschiedet sich diese Gattung von den anderen Thiaridae durch ein kurzes, anteriores Osphradium, ein Mitteldarm mit einem flach ausgeprägten Blinddarm sowie einen großen „accessory pad“ (zusätzlichem Ballen), der auf dem „glandular pad“ (Drüsenballen) aufsitzt. Anatomische, molekulare und statistische Untersuchungen zeigen, dass die angenommene Anzahl von ehemals fünf kubanischen Arten innerhalb der Gattung auf zwei, evtl. sogar nur auf eine einzige reduziert werden muss: die stark polymorphe, dennoch monophyletische Hemisinus cubanianus und H. martorelli, deren Status aufgrund des fehlenden Typmaterials fragwürdig bleibt. Aufgrund der phylogenetischen Analysen ist es weiterhin wahrscheinlich, dass die bislang Cubaedomus zugeordnete kubanische Art auch Hemisinus angehört. Diese Analysen zeigen weiterhin, dass die jamaikanische Art H. lineolatus eng mit den Thiaridae aus dem nördlichen Südamerika verwandt ist und eine basale Position gegenüber der kubanischen Art H. cubanianus einnimmt, was auch durch die anatomischen Untersuchungen bestätigt wird. Aufgrund der Ähnlichkeiten von anatomischen und morphologischen Merkmalen zwischen Hemisinus und anderen afrikanischen Familien, die den Thiaridae nahe stehen, lässt sich vermuten, dass diese einen gemeinsamen Vorfahren hatten. Dies wird zusätzlich durch die Topologie der phylogenetischen Analyse unterstützt. Die biogeografischen Untersuchungen der geologischen Daten und der Verbreitungsmuster der Hemisinus-Fauna der Großen Antillen deuten auf einen kontinentalen Ursprung mit einer anschließenden Kolonialisierung und Ausbreitung auf den Inseln hin. Bei den Pachychilidae Troschel, 1858 ist die Situation komplexer. Trotz der weiten Verbreitung dieser Familie in den Neotropen fehlte bislang eine umfassende taxonomische und systematische Revision dieser Gruppe, die hiermit vorgelegt wird. In Mittelamerika gibt es fünf verschiedene Gattungen und mehr als einhundert beschriebene Arten, die zudem meistens für relativ kleine Gebiete beschrieben worden sind, wie etwa einzelne Flüsse oder Flusssysteme. Obwohl die Pachychilidae von Mittel- bis Südamerika und im karibischen Raum vorkommen, beschränkt sich meine Arbeit allein auf die mittelamerikanischen Vertreter, da die Anzahl der Arten mit unklarem taxonomischen Status in dieser Region besonders hoch ist. Die kubanischern Pachychiliden wurden jedoch in die genetischen und biogeografischen Analysen miteinbezogen. Aufgrund des systematischen Durcheinanders und des schlechten Verständnisses der Beziehungen innerhalb der mittelamerikanischen und kubanischen Pachychilidae, wurden anatomische Untersuchungen des Gehäuses und der Radula sowie phylogenetische Analysen von zwei mitochondrialen Genen (COI, 16S) durchgeführt. Dabei wurden feine morphologische Variationen der Schale und der Radula zwischen den meisten Arten gefunden, die jedoch homoplasisch und somit wenig geeignete taxonomische Merkmale sind, um die neotropischen Pachychiliden-Arten unterscheiden zu können. Der Besitz von glatten Gehäusen ist ein konvergentes Merkmal, welches mindestens zweimal in der mittleramerikanischen Region entstanden ist, während die skulptierte Schale eine Synapomorphie ist. Ferner kann ich zeigen, dass die Pachychiliden-Arten aus Zentral- Mexiko sowie die kubanische Pachychilus und P. vallesensis in andere Gattungen als Pachychilus gehören. Die molekularen Untersuchungen zeigen darüber hinaus, dass die mittelamerikanischen und kubanischen Pachychilidae monophyletisch sind. Außerdem konnte ich die Existenz von verschiedenen, klar abgegrenzten Clades aufdecken , die (wahrscheinlich endemische) Populationen aus einzelnen Flüssen umfassen . Die wahre Diversität in dieser Familie wird aufgrund der großen Ähnlichkeit in der Schalen-Morphologie mit Sicherheit unterschätzt, wobei die meisten der beschriebenen Arten aus Mexiko und Guatemala als gültig betrachtet werden sollten. Zukünftige Revisionen werden höchstwahrscheinlich weitere, bislang noch unbekannte Arten identifizieren. Geologische, biogeografische und molekulare Daten deuten auf einen vikarianten Ursprung der kubanischen Pachychilidae sowie auf einen dispersiven Ursprung der kontinentalen Mitglieder der Familie hin, der von einem Vorfahren aus Zentral-Guatemala ausging. Abstract In this work I reviewed the systematics of two distinct lineages of Mesoamerica and Caribbean freshwater snails within the gastropod superfamily Cerithioidea: Thiaridae Gill, 1871 and Pachychilidae Troschel, 1858. Through comparative analyses of morphological, anatomical and molecular genetic data, I found that the family Thiaridae is represented by the genus Hemisinus in the Greater Antilles. Regarding Pachychilidae, the identity of Mesoamerican and Cuban species was reviewed. I also determined relationship patterns between them and their relatives from other tropical countries. Additionally, geographical range reconstructions of the members of both families were made based on the information available in museum collections and from field trips. Concerning the Neotropical Thiaridae, I redescribed all Hemisinus species from the Greater Antilles including the type, which is Hemisinus lineolatus from Jamaica. The genus is represented in the region by one endemic species in Jamaica (Hemisinus lineolatus), and probably two endemic species in Cuba (H. cubanianus and Cubaedomus brevis). The diagnostic characters that separate Hemisinus from the rest of the thiarid genera are a short and anterior osphradium, and a midgut with shallow caecum and large accessory pad. Anatomical, molecular and statistical analyses showed that the five former Cuban species assigned to the genus Hemisinus are reduced to two: the monophyletic H. cubanianus, possessing a highly polymorphic shell, and H. martorelli, the specific status of which must remain until its type material has been studied. Phylogenetic analyses showed that the Jamaican H. lineolatus has a basal position with respect to the Cuban H. cubanianus, which is also confirmed by anatomical studies. In addition, H. lineolatus is closely related to the northern South American Thiaridae. Moreover, according to the phylogenetic analyses, it is highly probable that the Cuban Cubaedomus is a member of Hemisinus. Based on the present findings I suggest that the sharing of anatomical and morphological features of Hemisinus with African families other than the Thiaridae could be indicative of common ancestry with an Oriental freshwater lineage. This is also corroborated by the topology of the phylogenetic analysis. Concerning the biogeography of the group, geological data and geographic distribution of the Greater Antilles Hemisinus fauna point to a continental origin with subsequent dispersal to the Islands. In the Pachychilidae Troschel, 1858, the situation is more complex. The family is widely distributed in the Neotropical region and a comprehensive taxonomic and systematic revision of this group was lacking. In Mesoamerica, five different genera and more than one hundred species have been described, some of which are restricted to rather small areas such as single rivers or river
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